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Breeding Variation in Female Kakapo (Strigops Habroptilus) on Codfish Island in a Year of Low Food Supply

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Breeding Variation in Female Kakapo (Strigops Habroptilus) on Codfish Island in a Year of Low Food Supply 64 AvailableNew on-lineZealand at: Journal http://www.newzealandecology.org/nzje/ of Ecology, Vol. 36, No. 1, 2012 Breeding variation in female kakapo (Strigops habroptilus) on Codfish Island in a year of low food supply Joanna Whitehead1*, Brad Case, Kerry-Jayne Wilson2 and Laura Molles Department of Ecology, Faculty of Agriculture and Life Sciences, Lincoln University, PO Box 84, Lincoln, New Zealand 1Present address: Department of Conservation, PO Box 29, Te Anau 9600, New Zealand 2Present address: PO Box 70, Charleston 7865, West Coast, New Zealand *Author for correspondence (Email: [email protected]) Published on-line: 30 August 2011 Abstract: We investigated why some mature females of New Zealand’s critically endangered parrot, the kakapo (Strigops habroptilus), did not attempt to breed during the 2005 breeding season on Codfish Island. At a population level, the initiation of kakapo breeding appears to correspond with years of mast fruiting of rimu (Dacrydium cupressinum) trees, with the proportion of females that breed each season dependent on the quantity of rimu fruit available. This research investigates possible links between habitat quality within individual home ranges and the breeding status of adult females during 2005, when the abundance of available rimu fruit was low. We assessed the importance of both home range size and habitat characteristics in determining breeding attempts. Foraging home ranges were characterised using radio-tracking and triangulation techniques. The relative importance of habitat variables in optimal breeding habitat was assessed using ecological niche factor analysis. Our results show that female kakapo breeding in 2005 had, on average, home ranges twice the size of those females that did not breed that season and the ranges contained a significantly greater quantity of mature rimu forest. Multivariate analysis illustrates female kakapo were effectively partitioning available habitat, as breeders’ foraging locations were positively correlated with high-abundance rimu forest with a tall canopy, described as optimal breeding habitat. In contrast non-breeders’ locations were weakly correlated with short forest containing little or no mature rimu forest. To maximise reproductive output each breeding season, conservation managers need to ensure that all breeding-aged females occupy optimal breeding habitat on Codfish Island. Removal to other islands of kakapo not required in the breeding population may enable females to increase their home range size and occupy better breeding habitat. Keywords: ecological niche factor analysis; habitat selection; habitat quality; home range size; radio tracking; reproduction; rimu Introduction either the patchiness of the fruit crops that trigger breeding or differences in female condition (Eason et al. 2006). This Despite having an annual gonadal cycle (Cockrem & Rounce research investigates the hypothesis that variation in breeding 1995; Cockrem 2006), kākāpō (Strigops habroptilus) breed attempts between females in low fruit years is associated with only once every 2–5 years on Codfish Island, the location of the the quality and quantity of critical food resources available entire population of breeding-aged females, in response to the within their individual foraging ranges. mast fruiting of rimu (Dacrydium cupressinum) trees (Elliott To test this hypothesis we estimate foraging home ranges et al. 2001; Eason et al. 2006). The recovery of this critically of adult female kākāpō on Codfish Island in a non-breeding endangered parrot is slowed even further by considerable year (2006) and compare home range characteristics between variation in the number of females that nest in some rimu- females that did and did not breed in the previous (2005) fruiting years. On a population level, breeding in this lek summer. We predict that individual female kākāpō that bred will species (Merton et al. 1984) appears to be correlated with the have larger foraging home ranges, since having a larger home quantity of rimu fruit available, with most females nesting in range should increase the likelihood that an individual will be years of high rimu fruit production (Elliot et al. 2006; Table 1). able to access more abundant, higher quality resources required However, in low rimu fruiting years only some female kākāpō for breeding (Millspaugh & Marzluff 2001). We also predict in the population nest (Elliot et al. 2006), causing concern for that there will be distinct differences in vegetation composition conservation managers as there is no explanation for what and environmental conditions within the home ranges of might be preventing some females from breeding. breeding compared with non-breeding individuals, reflecting Hypotheses advanced to explain what triggers kākāpō to differences in habitat quality and resource availability. breed all focus on the females receiving cues that there will Specifically we investigate whether (1) foraging home be sufficient ripe rimu fruit to raise their young. Whether range sizes or vegetation selection differ between females that these triggers are cognitive (Lignon 1974) or nutritional bred the previous summer and those that did not, (2) tenure (Powlesland et al. 1992), or mediated via weight gain (Harper on the island correlates with home range size or nesting, et al. 2006) or hormonal stimulation (Cockrem 2006; Fidler (3) the prevalence of vegetation types with a high rimu et al. 2008) remains unresolved. In a given breeding year, component inside foraging ranges differs between breeding variation in the proportion of females that breed may reflect and non-breeding females, and (4) the relative importance of New Zealand Journal of Ecology (2012) 36(1): 64-74 © New Zealand Ecological Society. Whitehead et al.: Breeding variation in female kakapo 65 Table 1. Breeding years on Codfish Island (since the first a range of habitat variables in characterising the ecological translocation of kākāpō in 1987) showing the level of the niche required for females to breed in a low-rimu-mast year. rimu mast, percentage of rimu branches bearing fruit and The results of these analyses assist us in predicting optimal the number of breeding-aged females that nested (Kakapo breeding habitat for female kākāpō on Codfish Island, thus Recovery Programme, unpubl. data) compared with the providing an additional tool for managing this critically total number on the island. The breeding year of 1992 is endangered species. not included as rimu mast and breeding attempts were not accurately recorded. ____________________________________________________________________________ Breeding Rimu Rimu bearing No. of breeding-aged Methods year mast fruit females* that nested level (%) (total no. on island) Codfish Island / Whenuahou (1475 ha) is situated 3 km off the ____________________________________________________________________________ north-west coast of Stewart Island in southern New Zealand 1997 Low 14 6 (10) (46°46.07′ S, 167°37.23′ E; Fig. 1). Habitat on the island is 2002 High 35 20 (21) dominated by mixed-podocarp forest with pakahi scrub in 2005 Low 11 10 (21) elevated areas and dense scrub around coastal margins (Meurk & Wilson 1989). At the time of this research in 2006, Codfish 2008 Low 13 5 (38) Island was home to the only known breeding population of 2009 High 39 27 (38) ____________________________________________________________________________ kākāpō as all 21 adult females of breeding age (9 years or *Note breeding age was defined as 9 years or older prior to 2008 older; Eason et al. 2006) resided there, along with 20 adult (Eason et al. 2006), and as 6 years or older from the 2008 breeding males and 13 juveniles. Our research involved 18 of the 21 season onwards as 6-year-old females nested that year (Moorhouse breeding-aged females; one female was excluded from the 2009). If a breeding age of 6 years or older had been used for the study because she was hand-reared and had formed attachments earlier years there would have been 26 breeding-aged females in to staff and two others were excluded because they were in 2005, but no change to the 2002 or 1997 figures. inaccessible locations on the island. We assigned a breeding status (i.e. breeder or non-breeder) to each of the remaining 18 adult females, based on the birds’ breeding status from the previous year (2005), which was the most recent breeding year (Table 1). Stewart Island Figure 1. Codfish Island / Whenuahou, located 3 km off the north-west coast of Stewart Island, New Zealand (NZMS260 1:50 000 scale map series provided by Land Information New Zealand). 66 New Zealand Journal of Ecology, Vol. 36, No. 1, 2012 Radio tracking of foraging locations 1989; Harris et al. 1990). We used a fixed kernel and the least- All kākāpō on the island are fitted with back-mounted radio squares cross-validation technique (Seaman & Powell 1996). transmitters, for management purposes. Foraging locations Home ranges were estimated using the software Ranges6 were estimated for each of the 18 female kākāpō between v.1.2 (Kenward et al. 2003). We used a regression model to 15 March and 30 May 2006, using standard radio-tracking determine whether the number of locations used to estimate techniques (Clout 2006). We used TR4 radio receivers a foraging range influenced its size. (Teleonics, AZ, USA) and three-element Yagi hand-held aerials We analysed the influence of 2006 foraging home range fitted with sighting compasses (Sirtrack, Havelock North, NZ) size on breeding status, which was determined using
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